Botanica Marina Vol. 45, 2002, pp.


© 2002 by Walter de Gruyter · Berlin · New York

A Revision of Amansia glomerata C. Agardh, Amansia rhodantha (Harvey) J. Agardh and Melanamansia glomerata (C. Agardh) R. E. Norris (Rhodophyta: Rhodomelaceae)
A. D. R. N’Yeurt
Jeune Equipe Terre Océan, Université de Polynésie Française, BP 6570, Faa’a Aéroport, Tahiti 98702, French Polynesia,

Species of Amansia and Melanamansia from the tropical Pacific region have been re-examined, and compared with material from the Indian Ocean, Japan, Taiwan and the Philippines. All specimens examined from the Pacific region (except Hawaii and some from New Caledonia and Fiji) lack pseudo-pericentral cells, thus placing them in Amansia. Melanamansia glomerata is only reported from Hawaii, New Caledonia, Fiji, the Philippines,Vietnam, Malaysia and for the first time in the Indian Ocean from Kenya.Type material of Amansia rhodantha (Harvey) J. Agardh from Mauritius was compared and found not significantly different from the Pacific Amansia material (including type and isotype material of Amansia paloloensis South et Skelton from Samoa, which is reduced to synonymy), thus extending the distribution range of A. rhodantha as a pantropical entity.

The genus Amansia was erected by Lamouroux (1809: 322) and is placed in the tribe Amansieae of the family Rhodomelaceae (Hommersand 1963: 335). The tribe is characterised by dorsi-ventral, erect thalli arising in tufts from a common basal disc. Most genera have five pericentral cells around the axial cell (Figs 1, 2) except Kuetzingia Sonder and Protokuetzingia Falkenberg, which have six pericentral cells (Kylin 1956). The two first-formed pericentral cells lie on the dorsal side of the characteristically inrolled apex of the mature lanceolate blades. In Amansia the dorsal and dorsi-lateral pericentral cells further divide to form immediately behind the apex a distromatic wing or ala, whose cells are arranged in alternate layers, with secondary lateral pit-connections (Fritsch 1945: 570). Recently, Norris (1988b, 1995) re-examined species of Amansia in the Indian Ocean and Hawaii, and discovered that some members of this genus differed consistently by having the first and second pericentral cells each producing a lateral pseudo-pericentral cell lying adjacent, but not connected to, the axial cell (Norris 1988b: 214; Fig. 3). Species previously ascribed to Amansia that had these pseudo-pericentral cells were transferred to a new genus (Melanamansia R. E. Norris 1988b: 217). Notably, Hawaiian plants previously ascribed to the common tropical species Amansia glomerata C. Agardh (whose type locality is Hawaii) were transferred to Melanamansia glomerata (C. Agardh) R. E. Norris, while Indian Ocean ‘Amansia glomerata’ were put under the earlier name Amansia rhodantha (Harvey) J. Agardh based

on their lack of pseudo-pericentral cells (Norris 1995). The genus Amansia currently consists of four species, the Caribbean A. multifida Lamouroux (the type of the generic name), A. loriformis R. E. Norris and A. rhodantha (Harvey) J. Agardh from the Indian Ocean and Malaysia, and A. paloloensis South et Skelton from Samoa. Melanamansia currently consists of twelve species, M. daemelii (Sonder) R. E. Norris, M. glomerata (C. Agardh) R. E. Norris, M. japonica (Holmes) R. E. Norris, M. mamillaris (Lamouroux ex C. Agardh) R. E. Norris, M. pinnatifida (Harvey) R. E. Norris, M. pumila (Sonder) R. E. Norris, M. scalpellata (Tanaka) R. E. Norris, M. seagriefii R. E. Norris, M. serrata (Harvey) R. E. Norris and M. mitsuii (Tanaka) Yoshida. Following these revisions by R.E.Norris,several authors (Yoshida et al. 1995,N’Yeurt et al. 1996,Payri and N’Yeurt 1997, Millar 1999, Millar et al. 1999) made the nomenclatural change from Amansia glomerata to Melanamansia glomerata in their checklists and herbarium collections from the tropical Pacific, but until recently no detailed studies verifying these changes as correct have been done. In 1995, samples of ‘Amansia glomerata’ from Rotuma Island (north of Fiji) were sent to Dr R.E.Norris,who confirmed (pers. com.) that pseudo-pericentral cells were present, and on the basis of this assumed finding the name Melanamansia glomerata was used in later publications (N’Yeurt 1996,N’Yeurt et al. 1996).Later,in the course of on-going investigations on the Samoan algal flora, South and Skelton (1999) found a species of Amansia lacking pseudo-pericentral cells, which they described as new (A. paloloensis South et Skelton), while in Malaysia and Vietnam (Masuda et al. 2000, Masuda

in EPHE). Sakai and S. 1989. showing intrusion of the first derivative of the first-formed pericentral cell into the axial ring (Suva. Figs 1–3. M. A. leg. Tahiti. 1997. Skelton and Dave [SUVA-A 1565 (holotype). Fiji. Punaauia. Rintels (SUVA-A 3891. 1999. Material and Methods Specimens were collected by reef-walking. – Samoa: Palolo Deep Reserve. Kagoshima: 9 August 1959. 3893. M.‘Al’ and ‘CG’ refer to specimens housed at IRD.) recent studies found both Melanamansia glomerata and Amansia rhodantha (Harvey) J. com. it was decided to revise records in the light of recent findings. otherwise as in Holmgren et al. deep algal rim. 1998. N’Yeurt (UPF 584). E. pers. 3892. – Taiwan: Anon. 1. P. 5480L (cystocarpic). 2. Afiti and D. D. A. 2 Jul. ps = pseudo-pericentral cells). Skelton. cross section of the lectotype from the Hawaiian Islands (LD 42600) showing the first two dorsal pericentral cells each developing a pseudo-pericentral cell (ps) that lies adjacent to the axial cell. leg. leg. 1998. . Tahiti. N’Yeurt (SUVA-A 5128). Tombleson and T. A. and a camera-lucida attachment was used to make anatomical drawings. A. in particular paying attention to important anatomical features such as the presence or absence of pseudo-pericentral cells and the number of segments between lateral veins of the blade (Norris 1995). – Cook Islands: Ngatangiia. Arrangement of pericentral cells in species of Amansia and Melanamansia (modified from Norris 1988b. Photomicrographs were obtained using an Olympus BX-50 compound microscope. 1998. 3898. A. (1981). 1–5 = order of formation of pericentral cells. Results Representative material examined Amansia rhodantha – Japan: Hanasado Beach. 1998. Prepared slides were examined using a Zeiss compound microscope. 9 Feb. P. Skelton and Dave (SUVA-A 1567). SUVA-A 1566 (isotype). leg. leg. Fig. 19 Mar. 11 Jul. N’Yeurt In order to clarify the taxonomy of species hitherto ascribed to Melanamansia glomerata. Amansia rhodantha (Beqa. leg. 3897). P. leg. 3896. leg. 2 Jul. SUVA-A 5438L). 25 October 1984 (SAP 046978). P. and stored in 5% buffered Formalin in seawater prior to shipment to the laboratory. Amansia rhodantha and Amansia glomerata.. Melanamansia glomerata. Sections were made using a freezing microtome. Okinoerabujima Island (60 km north of Okinawa). – French Polynesia: Moorea. Furusato. Faletose (SUVA-A 3894. Y. a = axial cell. SUVA-A 2402 (Lectotype) and liquid-preserved isotypes 5479L (tetrasporic). Baba (SAP 056160). Enomoto (SAP). leg. Fig. 5481L (spermatangial)]. 3895.Agardh to co-exist in the flora. R. A. U. 18 Sep. 6 Jun. 1984. M. N’Yeurt (UPF 347). Fig. Amansia rhodantha. 13 May 1982. C. Okinoerabujima Island (60 km north of Okinawa) 27 Sept. Fiji. Rarotonga.232 A. Payri (8/84. Nouméa. Most recently. Okidomari. 1998. SUVA-A 5436L). snorkelling or SCUBA diving. Tanegashima Island. 3. New Caledonia. leg. Palolo Deep. Macrophotography was done with a Nikon F2A in conjunction with Kodak Plus-X pan 125 film. developed and printed in the laboratory. Skelton. leg. and stained with cotton blue in lactophenol or 1% acidified aniline blue in 60% clear corn syrup. a detailed morphological and taxonomic study on some selected members of the Amansieae (Wilson and Kraft 2000) echoed the opinion that the genus Amansia in Australia was in need of critical revision. A. Baba (SAP 056163). The results of these investigations are described below. 3899). Taharaa.

leg. leg. G. 28 Nov. Hildebrandt (L 0108600). 84 (L 0108603). leg. R. 1973. B. 1834). leg. leg. Richer de Forges (AL 288d-CG 77a).Palawan. Isaac (L 0108594). paloloensis and Melanamansia glomerata specimens examined in this study. H.23 May 1978. 25 September 1999. Jul. Garrigue (CG 77c. Oahu. 1982.leg. leg. Nanuyalevu. n. – Philippines:Gao-oa. 24 Nov. leg.1 22. Bahati. – New Caledonia: Lajeu Sudouest. Cannon (SAP 052062).Solong-On. 14 Feb. 1995. SIPHILEXP-78 (US 088532). 84. 10 May 1978. 29 May 1978. Negros Oriental. SIPHILEXP-78 (US 013760). 17 Mar. Viti Levu. Ballou (SUVA-A 5438L). Table I presents the average results of morphological measurements carried out on Amansia rhodantha.Siaton. n. Farlow (L 0108597). G. Huisman (MURU. Yasawa Islands. Ningaloo. 26 Nov. – Indonesia: Cape Batu Kerapo.6 3. Skelton and G. S. Vatualailai. Species Thallus height (cm) Blade width (mm) Blade length (mm) Serrations (1 –3)* Cortication (1 –3)* Rosettes (1 –3)* Amansia rhodantha Amansia paloloensis Melanamansia glomerata 39. – Lord Howe Island: 18 Aug. leg. 16 Nov. R127).3 2. Negros Oriental. G. leg. 347). Littler (US 163490). Potipot Is. N’Yeurt (SUVA-A 5437L). leg. South (SUVA-A 2460). in TCD (Telfair. – Fiji: ‘Feejee’.Dumaguete City. Beqa Lagoon. leg. Yasawa Group. Kasahara (SUVA-A 1013. 1994 (SUVA-A 797).). Hapmak. leg. Wilkes Exploring Expedition (US 04057a). J. Amansia rhodantha: n. 1982. C.Siquijor Is. P. Isaac (L 0108595).11 Feb. leg. Chesterfield.9 18. 1838–42. 3 and 5 March 1992. 10 July 1992.. Figs 4–10 . Kauai. Kipukai. B79). 13 July 1972.W. leg.7 54. Huisman (CWC 1997. SIPHILEXP-78 (US 088531). 1979. 21 Sep. leg. 1980.. Lectotype. H. R. 1819. Snellius-II (L 0108601). leg. leg. P. leg. fide G. leg. Huisman (MURU. A. Silva et al. leg. Garrigue (CH1).d. JH240). Sumbawa.Malo. Garrigue (SUVAA 5436L). N’Yeurt (SUVA-A 346. 1876. 1980.. – Australia: Heron Island QLD. D. Viti Levu. 1983. anon. Barrow Island. 1995. Agardh 1841: 26. S. Taxonomic results Amansia rhodantha (Harvey) J. 1876. Magnesia Virae. W. – Fiji: ‘Feejee’. M. Agardh). leg. W. 07 Oct. leg. J. leg. Bay of Sanggar. Masuda et al. Villeneuve (SUVA-A 1737). A. H. leg.14 May 1979. Namada. 6 Jun 1978. 15 Oct. Plongeurs IRD (AL 288a. E. 1981. leg. Villeneuve (SUVA-A 1774). M. Mauritius. leg. Jul. W. 1985. AL 288b).Negros Oriental. A. 3 Jun 1978. leg. (MURU. leg. M. Barrow Island. Carlson (SUVA-A 281). 3 Feb. Hildebrandt (L 0108596. n. Rukuruku Village.2 53.Negros Oriental.. leg. 17 Jul. E. A. South (SUVA-A 5544. R. Candelaria Zambales. 1995. SIPHILEXP-78 (US 088534). Gaudichaud (‘Amansia glomerata’. 1965. Tantabiddi. Hakula. 18 March 1992. Average values of selected characters for Amansia paloloensis. 84. leg. L 0108598). Suva Reef. Unia. Melanamansia glomerata – Hawaiian Islands: Hawaiian Islands. A.7 2 2 1 2 2 2 2 2 2 * As measured on an index of (1 = little) to (3 = pronounced) – Rotuma: Hoféa. A. leg.Canipo Is.. Naukathura Island.5 3. J. Oahu.. SIPHILEXP-78 (US 088530). – Mozambique: Ponta Abril. leg. C. South (SUVA-A 5431). Norris 1988b: 211. M. 2000: 188. Amansia rhodantha and Melanamansia glomerata surveyed in this study.Revision of some Amansia and Melanamansia species 233 Table I. Lopta. 1985. J. 1979. R. B. 16 March 1991. Inhaca Peninsula. leg. 3 and 15 Oct. CG 77b). 1996: 472. see text). Ovalau. Daida (US 086755). SIPHILEXP-79 (US 088533). Keats. Cordero and Tuangpalan (US 091395). figs 1–11 (as Amansia glomerata C. – Kenya: Mombassa. 1067).d. leg. B262).Moreland (US 50851). Basay.Calindagan Reef.d. 1984. leg. 10 Aug. 27 Dec. C.leg. Lawai Kai. 5 June 1967.. Dumaguete. Snellius-II (L 0108602). J. – Mauritius: Delesseria rhodantha Harvey (holotype) (= Amansia rhodantha): Cap Malheureux. 14 March 1998. figs 37–42. 06 Sept. n.. 1984. Oahu. A. leg. Montebello Islands. M de Robillard (L 0108599). (US 02890). Ti Tree.Cuyo. – Philippines: Actin. C. Cordero and Tuangpalan (US 091394). P. 9 Sept. B. leg. J. Viti Levu. Banilad. 25 Feb. leg. E. Suva Barrier Reef.d. Magruder (US 086459). 1954. R.Ilocos Norte. Cribb (SAP 051714). 30 Jun. 21/22-Sept. J.9 11. c. leg. 24 April 1996. A. leg. South in lit. 16 Oct.d. Hanauma Bay Beach Park. 1988. leg. – New Caledonia: Lajeu Sudouest. Huisman. leg. leg. G. Catanduanes. – Kenya: Mombassa. Jurien Bay WA. leg.P. NR 248).Pagodpud. W. 1997. Kauai. 1838–42. LD 42600 / 0772). Wilkes Exploring Expedition (US 04057b) (doubtful record. 1069). leg. Salayer. Huisman (MURU. Eagles Nest. Tangangge. leg. Kasahara (SUVA-A 1012. Diani Beach.

Mauritius. and a pair of budding branchlets from the midrib in the lower half. Fig.234 A.5 mm). Surface view of segment cells of blade alae. Indian Ocean. Figs 4. 6. 8. from Rotuma Island. . Amansia rhodantha. US 02890 (scale bars = 3 mm). Secondary branchlet. in TCD (scale bar = 25 µm). Type material of Delesseria rhodantha Harvey. Type material of Delesseria rhodantha Harvey. in TCD (scale bar = 200 µm). 5. South Pacific (scale bar = 10 µm). Habit of mature plant from the type locality. showing both lax and dense growth forms. Fig. SUVA-A 5437L. N’Yeurt Figs 4–10. showing axial cell (a) surrounded by five pericentral cells. Fig. D. Axial structure of blade. 10. Fig. 9. 7. showing marginal serrations. Detail of tetrasporangial stichidia on lateral endogenous branch. R. in TCD (scale bar = 2. Rehydrated type material of Delesseria rhodantha Harvey.

showing both lax branching and the presence of rosettes (arrowheads) (scale bar = 4 mm).Revision of some Amansia and Melanamansia species 235 Figs 11–16. Tetrasporangial stichidia of Amansia rhodantha from Beqa. Segment cells of blade alae in surface view. Detail of the holotype (SUVA-A 1565). Note marginal cystocarps (arrowheads). SUVA-A 3892 (scale bar = 12 mm). SUVA-A 1565 (scale bar = 25 µm). Fiji. Fig. Holotype of Amansia paloloensis. Fig. with numerous rosette axes. SUVA-A 5436L (scale bar = 50 µm). .5 mm). showing lateral endogenous branches and midrib. SUVA-A 3892 (scale bar = 2. SUVA-A 5479L (scale bar = 1 mm). Compare with Fig. Habit of mature plant. 11. Fig. 14. Detail of blades. 12. 16. Detail of morphologically distinct mature rosette and corticated axis. Fig. 4. 15. Fig. 13. Amansia paloloensis from Samoa. Fig.

and the apex is characteristically in-rolled. Vietnam. Melanamansia glomerata. Agardh) R. to 0. and two displaced rows of regularly arranged hexagonal alar cells 20–29 × 92–129 µm in surface view. Note that the stichidia is twice laterally branched. Plant from Potipot Island. while cystocarps 700–1000 µm in diameter occur usually singly and terminally on endogenous branches. 1999: 573. 1987: 61. US 091395 (scale bar = 5 mm). showing smooth margins and elongate shape. 20. 22. Carposporangia are lanceolate. Millar 1999: 523 (unverified record). Cribb 1983: 106. Okamura 1900: 71. Fiji. Blade margins are smooth. figs 19–21. N’Yeurt et al. Norris 1995: 66) is usually 4. Fig. plate CXXVI. Fig. 4B. Ultimate branchlets are ecorticate. L 0108598 (scale bar = 700 µm). e–i. 100–112 µm in diameter. Tetrasporangial stichidia are pinnately arranged on either side of blade margins. 185. Branching is lax to profuse and usually forms deep-red leafy rosettes of secondary branchlets when fully mature. 18. pl. Yoshida et al. 118C–J. Melanamansia glomerata (C. Coppejans and Millar 2000: 333 (unverified record). 21. 23.5 µm in diameter that lies adjacent to the axial cell (Figs 3. Deciduous. containing up to 10 Figs 17–24. 1996: 87. Blade of plant from Lajeu. Fig. or with regular serrations (endogenous branches). denuded below. and are crisp and lanceolate. 19. New Caledonia. D. CG77a (scale bar = 10 µm).8–14. New Caledonia. figs 1–11. Falkenberg 1901: 416. 77–110 µm thick. 9). XXV. fig. with the first two dorsal pericentral cells each developing a smaller pseudo-pericentral cell 13. up to 100 (on average 40) mm high. Weber-van Bosse 1923: 369. Trichoblasts are sparse. 0. New Caledonia. Payri and Meinesz 1985: 511. figs 2–14 (type locality: Palolo Deep Marine Reserve. with more or less pronounced marginal serrations 1–4 mm long (which represent endogenous branches) always present. 17. The number of segments between alternating veins (determinate branches covered by the corticating pericentral cell derivatives. Fig. A central midrib is present. Individual tetrasporangia are spherical and tetrahedrally divided. 2. The stem of plants is thick and cartilaginous. 6. with a central axial cell 28–30 µm in diameter surrounded by five pericentral cells 36–40 µm in diameter (Figs 1. they are up to 4 cells long and non-vesiculate. The number of segments between alternating veins (determinate branches covered by the corticating pericentral cell derivatives. on average 53 mm high. East Africa. 1. 212a. 128. Philippines. pl. holotype in TCD. fig. Tropical Pacific Ocean. 31 fig. E. Agardh 1824: 194. 198. 6E (unverified record). Samoa). Detail of blade of plant from Mombassa. colourless and uniseriate dorsal trichoblasts up to 8 cells long develop on every second or third axial cell (or on every segment in determinate marginal branchlets). Norris 1995: 67. The central axial cell is 28–30 µm in diameter. Agardh 1824: 247. Silva et al. with a range of 1–5. becoming narrower and disappearing towards the apex. Plant from Unia. 1. with a range of 3–6. 85–110 µm thick. figs 1–3 (type locality: Cap Malheureux. A distinct midrib up to 1 mm wide is present in the lower half of ultimate branchlets. Japan. Garrigue and Tsuda 1988: 63. Abbott 1999: 404. South-East Asia. up to 35 mm long and 6 mm broad. Isaac and Isaac 1968: 24. Kenya. 197. 1995: 150. fig. Kylin 1956: 544. Mauritius.4. Distribution: Tropical Indian Ocean. Internal structure is cellular. Yamada and Tanaka 1938: 86. New Caledonia. N’Yeurt Basionym: Delesseria rhodantha Harvey 1834: 151–152. surrounded by five pericentral cells. figs 1–13. 436B. Kenya. The thallus is rose-red. Dublin). Millar et al. E. in up to 10 regularly disposed pairs. up to 750 µm long and 250 µm broad. stem-like and rigid in basal portions. Agardh) R. Fritsch 1945: 570. Known distribution: Hawaiian Islands. Lewis and Norris 1987: 22. fig. pl. 19). Jaasund 1976: 131. AL288b (scale bar = 5 mm). The main axis is irregularly branched. Tsuda and Wray 1977: 103. pl. Norris 1995: 67. Synonymy: Rytiphlaea rhodantha (Harvey) Decaisne 1842: 358. US 4057b (scale bar = 1 mm).5–0. Tetrasporangial stichidia of plant from Oahu. Plants are brownish-red. 20. 267. Fig.236 A.8 mm in diameter. 25–42 × 83–125 µm. Apia. Norris 1988b: 211. Amansia paloloensis South et Skelton 1999: 247. Procarps and curved spermatangial heads are 200–450 µm in diameter and are dorsal on the involucral tips of endogenous lateral branches. Plant from Mombassa. figs 14–29. figs 127. Axial structure of plant from Lajeu. Isaac 1956: 188. Blades of plant assumed to be from the Fiji Islands (see text). Fig. fig. L 0108598 (scale bar = 5 mm). Philippines. SUVA-A 5431 (scale bar = 50 µm). 24. showing pseudo-pericentral cells (arrowheads) surrounding (but not directly connected to) axial cell (a). Secondary blades develop as successive pairs on the dorsal side of the midrib. fig. Womersley and Bailey 1970: 336. . composed of two overlapping layers of elongate hexagonal cells in transverse rows. – Melanamansia glomerata (C. N’Yeurt 1996: 428. with in-rolled leaf apex. CG77a (scale bar = 1 mm). Norris 1995: 66) is on average 4. Marginal teeth are up to 1 mm long and 445 mm broad. Tetrasporangia occur in flattened. pl. pinnately arranged marginal stichidia up to 340 µm long and 290 µm in diameter. Fig. Figs 11–16 Misapplied names: Amansia glomerata C. Figs 17–24 Basionym: Amansia glomerata C. each cell is about 20–20 × 87–100 µm in surface view. Kenya. and are composed of lanceolate to ovate blades to 100 mm long and 3 mm wide. and are initially vesiculate. 7 mm broad. R. Payri and N’Yeurt 1997: 897. Fig.

and stained the herbarium mounting paper.Revision of some Amansia and Melanamansia species 237 pairs of tetrahedrally-divided tetrasporangia which are 50–100 µm in diameter. Spermatangial heads are curved and arranged in marginal clusters. In all cases. . procarps and cystocarps occur on endogenous marginal branches. the colour of the Melanamansia plants was noticeably brownish-red.

More importantly. the absence of blades organized in morphologically distinct tight axes around the parent axis. an additional field test to distinguish Amansia spp. 14) and corticated rosette axes (Fig. Smith (1979: 38–40) points out that the accounting and . with for instance A. Now that the Samoan species has been compared with genuine herbarium material of A. On the other hand. paloloensis showed no major difference in the range of characteristics among them. which is welldocumented and illustrated in the protologue of the species. 7) shows features similar to those seen in blades of South Pacific material previously ascribed to Amansia glomerata. is comparable to A. rhodantha from Western Australia (MURU B262. 3) which distinguishes that plant from A. paloloensis from the type locality. isotype and other collections of A. 23) but the plant has not been collected in Fiji waters since. the Philippines. Kenya. rhodantha. and a cross section of the blade clearly shows the five pericentral cells around the axial ring. The drawing of the holotype (South and Skelton 1999: 246. indicating that the morphology of the species is influenced perhaps by environmental factors. while Melanamansia becomes dark brown. rhodantha from the type collection and localities in Mauritius. Vidalia. The Wilkes collection consists of two plants. a comparison of material of A.).238 A. having tight rosettes as well as very lax. From this study. MURU 248). regularly alternate marginal serrations which are not as pronounced as in Amansia rhodantha and which are often absent. and absence of pseudopericentral cells. a single record for this genus exists (collected from ‘Feejee’ by the Charles Wilkes U. Kuetzingia. However. Mozambique and Western Australia. Agardh from Hawaii (LD 42600 / 0772) clearly shows the presence of paired dorsal pseudo-pericentral cells in a blade cross section (Fig. Melanamansia glomerata in the tropical Pacific Wilson and Kraft (2000: 366) commented that the generic segregation of Melanamansia from Amansia is probably not warranted. paloloensis from A. it appears to be conspecific with the latter species and thus extends the status of A. more coriaceous branchlets with less apparent midribs (this study). apparently because the brown pigments (presumably polyphenols) are less susceptible to the bleaching action of the preservative (as is the case in species of brown algae such as Sargassum and Turbinaria). and found a significant number of thalli with morphologically distinct primary corticated axes and relatively tight and lax rosettes of secondary blades on rosette axes (Figs 11. Fiji. Exploring Expedition. The reproductive and trichoblast anatomy of A. serrate branches and lax ‘rosettes’ of secondary blades issued from the weakly developed midrib. The present author has re-examined the holotype. material from the Tropical Pacific and Indian Oceans previously known as Amansia glomerata (or misidentified as Melanamansia glomerata. rhodantha as a pan-tropical entity. 12). fig. The status of Amansia paloloensis South et Skelton South and Skelton (1999) described a new species of Amansia from Samoa. including the presence of both tight and lax rosettes of secondary branches and alar cells on the primary axes. the Cook Islands and French Polynesia (Table I) with A. Japan. M. paloloensis. a re-examination of the lectotype of Amansia glomerata C. Fig. from Melanamansia glomerata was found: when preserved in 4% Formalin-seawater and exposed to ambient light for several days. between 1838 and 1842 (US 04057b. Rotuma. R. While there is no doubt as to the generic identity of the latter specimen. 2) indeed shows a plant with loriform. rhodantha as described and illustrated by Norris (1988b: 211). figs 1–4) indicate features quite similar to those found in the Amansia species considered in this study from localities in both the Indian Ocean and the tropical Pacific Ocean. length of the blades. degree of cortication of the midrib and position of the reproductive stichidia are equally quite variable in plants from the same and similar populations. thicker. and Ceramium. one of which (US 04057a) is Amansia rhodantha. rhodantha (South and Skelton 1999: 249). Characters such as the degree of rosette formation. while the other (US 04057b) is Melanamansia glomerata. and these authors considered the lack of highly corticated axes. Masuda pers. These plants also had blades with well-developed midribs in the lower half (Fig. generally smaller.13). plate CXXVI. D. In the Fiji Islands. loriform blades reminiscent of some specimens of A. N’Yeurt Discussion The holotype of Delesseria rhodantha Harvey An examination by the author of a rehydrated blade fragment of the holotype of Delesseria rhodantha Harvey in TCD (Fig. New Caledonia. since the presence of pseudo-pericentral cells is an inconsistent attribute in species of Neurymenia. Amansia tends to bleach to translucent white. paloloensis. Eastern Australia. and a weakly developed midrib as characters distinguishing A. as well as Amansia species from Taiwan. and trichoblasts lacking protective vesicles when young (Norris 1988b). rhodantha from the Indian Ocean and other localities.S. On the basis of these observations. when pseudo-pericentral cells were found to be absent) are considered conspecific with Amansia rhodantha. The habit and description of the holotype by Harvey (1834: 151. additional distinguishing characteristics of Melanamansia are a darkly coloured brown thallus which stains the mounting paper when dried (possibly linked to differences in chemical composition. com.

both on the same plant and within individuals in similar populations. when present). more corticated secondary and primary axes with tight rosettes. It would seem that perhaps shallow. thus placing them in Amansia. Herbarium and liquid-preserved samples of Amansia and Melanamansia species from the Pacific.Revision of some Amansia and Melanamansia species 239 recording of the Wilkes Expedition collections was not very satisfactory. which also had a consistently thicker thallus. perhaps an adaptation to the exposed habitats where the plants came from. both have an ala) and are arranged in ‘lax’ rosettes (South and Skelton 1999: 247. an examination of the holotype and liquid-preserved and pressed isotypes of A. 1) shows both the presence of tight and lax rosettes of secondary branches. However. the Philippines. and in some instances lacking altogether. type of endogenous branching. 5. degree of marginal serrations and thallus flattening. some specimens were virtually indistinguishable from specimens of A. Herbarium specimens of ‘Amansia glomerata’ from southern Japan housed in SAP (Hokkaido University) were examined. Payri and N’Yeurt 1997) were reexamined. pl. M. and found to consistently lack pseudo-pericentral cells and a similar observation was made on samples from Taiwan. pseudo-pericentral cells were universally absent. in New Caledonia and Kenya). and a main axis with alar cells not very distinct from secondary branches. and would seem to function as attachment haptera.g. the description of the species states that branches have the same shape as the main axis (i. CXXVI. 19). while younger plants have less cortication and more lax or absent rosettes of blades. com. more exposed conditions (such as on fringing reefs) favour tight rosette formation (which is more hydrodynamically resistant) while more lax arrangements are possible in deeper. the presence or absence of bladelets in rosettes. and is backed by other morphological and biochemical differences (Norris 1988b. environmental factors seem to play a role in determining the degree of rosette formation. There is a possibility it might have been mixed with collections from neighbouring localities such as the Solomon Islands. 13). Indonesian and Indian Ocean region were sectioned and examined for pseudo-pericentral cells. Wilson and Kraft 2000). in some cases. except the Hawaiian samples. which is a characteristic feature of both Amansia and Melanamansia (Norris 1988b) can vary from lax to tight. 41) similarly report the absence of pseudo-pericentral cells in A. M. rhodantha from Mauritius and Western Australia (compare Figs 4. fig. position of the reproductive axes and the distinctness of the axial ring in rosette axes (Norris 1988a. they were clearly apparent and about half the size of pericentral cells (Fig. and the presence of tight and lax rosettes of secondary branches on the same plant (Figs 11–13). older plants tend to have thicker. For instance. The author has unsuccessfully tried to obtain material from the latter locality. These ventral. Ecological studies would be needed to verify these hypotheses. while in Malaysia Masuda et al. irregularly dichotomous proliferations lack alar cells. calm waters (such as blue holes. Validity of current criteria to distinguish the Amansieae Characters that have proved useful in distinguishing species in the Amansieae include the presence or absence of pseudo-pericentral cells. 2). Indian Ocean Amansia rhodantha also lacked such cells.e. Also. pers. alternate marginal proliferations (which represent non-determinate or endogenous secondary branchlets) 2–5 mm long (Fig. and found to lack pseudo-pericentral cells. which was the main reason for separating the species from A. paloloensis. the vesiculate (in Amansia) or non-vesiculate (in Melanamansia) nature of trichoblasts. glomerata occurs in the same habitat as Amansia rhodantha (e. 1995). Fijian and French Polynesian specimens previously ascribed to Melanamansia (N’Yeurt et al. Proliferations from the ventral midrib (aside from secondary branchlets) were only found to be abundant in some of the Indonesian Amansia rhodantha specimens from Salayer (e. adding to the confusion between the two genera. All Amansia rhodantha specimens examined had fairly prominent. in the case of A. (2000: 188. Indeed. From an examination of the specimens at hand. and some samples from Kenya.g. it was observed that rosettes of blades along the secondary axes. rhodantha. Masuda. 1996. The presence or absence of these pseudo-pericentral cells hence appears to be a good generic criterion. The illustration by Harvey of the holotype of Delesseria rhodantha (1834. however it is clear that the degree of rosette formation and cortication of the axes is not a reliable taxonomic character at the species level in Amansia. glomerata occurs in the Solomon Islands. Melanamansia glomerata is unlikely to be confused with other Melanamansia species because of its unique combination of characteristics (Norris 1988b. rhodantha.). Fiji and New Caledonia (Table I). Generally. in the case of Amansia paloloensis). paloloensis revealed a wide range in the degree of rosette formations and cortication of the secondary axes and presence of alae on primary axes. fig. L 0108603). In Melanamansia glomerata these marginal serrations were much less pronounced (but more regular than in Amansia rhodantha. Norris (1988b: 211) reported that rosette axes in . fig. When pseudo-pericentral cells were present. In most instances. so it cannot be confirmed at this stage if M. However. 7). and hence the precise locality and date of this Melanamansia collection from Fiji is unknown. with the blades smooth and superficially reminiscent of those of some Sargassum species (Figs 22–24).

25. One feature which was noted in Melanamansia glomerata is that individual tetrasporangial stichidia are often laterally branched once or twice (Fig. Cells of the blade alae in both Amansia and Melanamansia are regularly elongate-hexagonal. the number ranged from 3 to 7. rhodantha and A. 16) and Melanamansia show no particular difference.3. paloloensis specimens studied. and occur in two overlapping and slightly offset layers radiating on either side of the midrib.240 A. Biogeographical map for Amansia rhodantha (circle) and Melanamansia glomerata (triangle). An examination of these elegant cells in species of Amansia (Figs 10.4 (Norris 1995). but the range is reportedly less than in Amansia rhodantha. which are in fact determinate branches covered by the pericentral cell derivatives. Hence the distinctness of the axial ring was not found a reliable taxonomic character among the species examined. This also varied according to the degree of rosette formation and age of the plant. There appears to be no notable difference in the range of thallus size. However. usually 3 or 4 with an average of 3. which was not seen in Amansia rhodantha (Figs 6. In Melanamansia it averages 4. paloloensis. which was also recorded by Norris (1988b) for Hawaiian material. Melanamansia glomerata from all localities studied have consistently smoother. R. 15). Tetrasporangial stichidia are basically similar in both Amansia rhodantha and Melanamansia glomerata. but can also occur apically. only being more or less surrounded by cortication. as the axial ring was still discernible in all sections made. D. as well as between A.). In A. as well as more corticated axes. At this stage there seems to be no largely significant difference in the number of segments between alternating veins in the specimens of A. N’Yeurt Amansia from Natal have an ill-defined axial row of cells. This feature was examined on a range of specimens of Amansia from the South Pacific and the Indian Ocean but no significant difference was seen in these specimens. B. smaller and narrower blades. Norris (1995: 66) counted the number of segment cells between these alternating veins for several species of Melanamansia and Amansia. cortication of axes and rosette formation between specimens of Amansia rhodantha studied. with most specimens having segment lengths of about 100 µm. From Table I. These differences reinforce the conspecificity of the Amansia rhodantha and A. rhodantha studied. . in elongate flattened and pinnate stichidia which are usually on lateral marginal branchlets. An additional potentially useful character is the number of segments between lateral ‘veins’ on the blades. rhodantha specimens observed in this study. and found them relatively constant. with the length of segments ranging from 80 to 120 µm. showing the currently known distribution. a number of preliminary conclusions can be made: A. Fig. Cover cells usually totally cover the developing sporangia. it was noted that in Melanamansia the range was more restricted. These can be easily detected in the unrolled apical portions of the blades by the developing trichoblast terminating each such alternating ‘vein’ (Norris 1995 and pers. blade dimensions. com. as well as the generic distinctness of Melanamansia glomerata. 20). in that they are typically rhodomelaceous with pairs of tetrasporangia in up to ten rows.

Washington).A. 1809. Skelton (SUVA – The University of the South Pacific). 443–457. Lund). Masuda. Die Gattungen der Rhodophyceen. W. J. Schofield. Scheltema & Holkema. Mar. Seven species of Rhodophyceae. Notice of a collection of algae. B. Dublin). Professor G. Volume II. E. Mr Barrett Brooks and Dr John Sims (US – Smithsonian Institution. E. N. Phil. Sapporo).). U. K. J. 1956. Rhodophyta. The Structure and Reproduction of the Algae. J. Bishop Museum Press. Agardh. 1983. sparse. Mus. Mar. com. Honolulu. (Hooker) 1: 147–157. Index Herbariorum. 1963. Intertidal Seaweeds in Tanzania. with descriptions of some new and little known species. P. Bot. Figure 25 presents a world distribution of these two genera as it is currently known. J. Bohn. Ms Susanna Riebe (LD – Botanical Museum. Isaac. University of Tromsø. East Africa Nat. Revised Key to the Species of Amansia and Melanamansia in the South Pacific – Presence of paired dorsal pericentral cells. Micronesica 21: 53–70. Dr R. 24 pls. com. Bot. frontispiece. xvi + 477 + [1] pp. 17: 297–380. Norris. S. Shimada. Leiden). Monographie 26. Berlin. African Bot. Keuken and E. Utrecht. H. First edition. The morphology and classification of some Ceramiaceae and Rhodomelaceae. 2000. Sci. Nat. K. Dr John Parnell (TCD – Trinity College. 1842. C. trichoblasts on every second or third axial cell. Kylin. pls cxxv. Soc. xiv + 939 pp. 173 + (2) pp. and A. – Absence of paired dorsal pericentral cells. F. T. 1981. Isaac. pigments reddish-brown. Isaac. Nouv. Rhodophyceae. Accepted 17 January 2002. and F. Fauna und Flora des Golfes von Neapel. 1824. R. flora and vegetation. In historiam algarum symbolae. A. Hooker by the late Mrs. et description de cinq nouveaux genres de cette famille. Sci. 458 figs. It is clear that both A. iii + 159 pp. xv + 673 pp.. E. ser. Observations sur la physiologie des algues marines. A history and annotated account of the benthic marine algae of Taiwan.A. Foreword. E.). Professor Willem Prud’homme van Reine (L – Rijksherbarium. 2000. M. Agardh. S. M. Tsuda. General account of the environment. Professor G.. Annal. Coppejans. The Herbaria of the World. Paris 1: 330–333. E. J. 2. Kato. 336 figs. xxxviii + 312 pp. M. Cambridge University Press. Mar. Professors Michio Masuda and Tadao Yoshida (SAP – Hokkaido University. Cal. Robin South is also thanked for his helpful comments on an earlier draft of this manuscript. Taxonomic notes on marine algae from Malaysia. Bull. Marine red algae from the North coast of Papua New Guinea.. E. Kawaguchi and S. J. Hommersand. Marine algae of Inhaca Island and of the Inhaca Peninsula. I. Phang. Kenya (this study). Norris is thanked for his helpful insights and suggestions for this research. Essais sur une classification des algues et des polypiers calcifères de Lamouroux. Phaeophyceae. F. 1945. I. J. Gleerups Förlag. Lamouroux. South pers. Handbook No 2. Harvey. A. trichoblasts non-vesiculate. 2 folded maps. Falkenberg. composed of up to 4 cells: Melanamansia glomerata. in the Mauritius. C. 71 pls. 1834. Cribb. Jaasund. 1956. New Caledonia and the Philippines (this study). (2000) confirm that the genus Amansia does exist in the Pacific Ocean. P. communicated to Dr. rhodantha and Melanamansia glomerata are more widely distributed than previously thought. Linnaea 15: 1–50. ments rose-red. 1999. 7th edition. 1901. Hist. M. K. W. Systema algarum.. Masuda pers. and J. Acknowledgements The author sincerely thanks the following persons and institutions for the generous loan of type material and specimens used in this study: Dr John Huisman (MURU – Murdoch University). V. Pub. Part I. E. G.Vietnam and Malaysia (M. Marine botany of the Kenya coast. Marine Algae of the Southern Great Barrier Reef. 1987. Bot. W.Revision of some Amansia and Melanamansia species 241 Conclusions The results of this study and those of South and Skelton (1999) and of Masuda et al. and Dr A. 43: 315–346. cxxvi. Dr Claude Payri (UPF – Tahiti). and possibly other localities which are not yet adequately investigated. Holmgren. 43: 181–190. and it appears that Melanamansia glomerata is so far restricted to Hawaii (Norris 1995). J. from “Cap Malheureux”. Garrigue. Robin South and Mr Posa A. Bot. W. Myxophyceae. Lord Howe Island (G. and each genus is consistently differentiated from the other by a number of anatomical and biochemical characters. pig- References Abbott.. Die Rhodomelaceen des Golfes von Neapel und der angrenzenden Meeres-Abschnitte. Marine Red Algae of the Hawaiian Islands. Charles Telfair. initially vesiculate and composed of up to 8 cells: Amansia rhodantha.. xvi + 754 pp. Contrib. and R. Fritsch. pls v–vii. . Nat. K. H. Lund.. 10 figs. Soc. Sci. 1976. Part 1. Millar for his generous assistance in obtaining copies of rare publications. 1968. Millar. Lewis. 1988. Catalogue of marine benthic algae from New Caledonia. 27: 7–28. K. Australian Coral Reef Society: Brisbane. H. Bot.W. 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